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Z O O L O G I E Contribution a L'etude Des Metamorphoses Chez Les Nudibranches

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Z O O L O G I E Contribution a L'etude Des Metamorphoses Chez Les Nudibranches ANNALES S0C1ÉTÉ f r a k, çasse ds MaUcolog¡£ SCIENCES NATURELLES 55' RUE ßUFFON P Á R j S - 5 6 ZOOLOGIE ET BIOLOGIE ANIMALE E XT FIA. I T CONTRIBUTION A L’ETUDE DES METAMORPHOSES CHEZ LES NUDIBRANCHES Par Jean TARDY (Laboratoire de Zoologie, Faculté des Sciences, 40, avenue du Recteur-Pineau, 86-Poitiers.) 1 2 e S érie — T om e X I I — Fascicule 3 — 1 9 7 0 MASSON & Cie, ÉDITEURS 120. BOULEVARD SAINT-GERMAIN. PARIS (VI*) Annales des Sciences Naturelles, Zoologie, Paris. 2441 5 12e Série, Tome XII, 1970, pp. 299-370. Vlaams Instituut voor de Zee CONTRIBUTION A L’ÉTUDE DES MÉTAMORPHOSES CHEZ LES NUDIBRANCHES(1) Par Jean TARDY (2) (Laboratoire de Zoologie, Faculté des Sciences, hO, avenue du Recteur-Pineau, 86-Poitiers.) ABSTRACT Contribution to the study of the metamorphosis in Nudi- branches. A careful study of the organogenesis ofÆolidiella alderi has been made. Other unknown ontogenic cases have been observed and compared with. Normal and teratological organogenesis, comparative anatomy lead to a discussion which particularly concerns phylogeny of Nudibranches. RÉSUMÉ A. — Le développement complet d’Aeolidiella, alderi étudié in vivo et à l’aide de coupes histologiques montre plus spéciale­ ment : I o La formation du système nerveux. 2" La torsion de 180°. 3° La réflexion du bourrelet palléal donnant le tégument dorsal et les papilles. 4° La réalisation de l’aspect limaciforme excluant toute possi­ bilité de détorsion. 5° La grande similitude d’organisation des véligères 1 des Nudibranches dont les œufs sont riches en vitellus. (1) Ce mémoire constitue la deuxième partie, dans sa version intégrale, d’une thèse de doctorat d’Etat (Sciences naturelles), C.N.R.S. n° 3287. (2) Avec la collaboration technique de Mmc C h e v a l i e r et MIle C h o p i n (C.N.R.S.). Je tiens à rendre hommage à l’ensemble du personnel du Laboratoire qui m’a rendu de multiples services au cours de ce travail. 300 JEAN TARDY B. — Comparaison avec les autres types de métamorphoses des Nudibranches : I o Aeolidiacea 7 à véligères planctotrophes : des véligères de Facelina coronata et d’une espèce indéterminée, à la métamor­ phose, montrent que la coquille croît de plus d’un tour durant leur vie pélagique. 2° Aeolidiacea 2. Chez toutes les véligères la portion du man­ teau qui délimite la cavité palléale a disparu à l’éclosion. La coquille larvaire ne peut donc croître pendant la phase péla­ gique. Le notum et les papilles naissent du plancher et des côtés de la cavité palléale. Les Aeolidiacea 2 sont plus évolués et la phase pélagique est plus courte que chez les Aeolidiacea i. 3° Pleurobranchacea : La morphogenèse deBerthella plumula est très semblable à celle observée chez les Nudibranches 1. C. — La discussion porte principalement sur : I o La torsion et sa signification adaptative. 2° La chronologie des processus de métamorphoses. 3° La tératologie, la morphogenèse naturelle et expérimentale chez les Nudibranches. 4" Le système nerveux. 5° Les conclusions phylogénétiques générales; l’évolution au sein des Nudibranches a): à partir de l’embryologie et b) de la tératologie et de l’anatomie comparée : les observations abou­ tissent à considérer les Nudibranches comme constituant un ordre naturel clairement défini. SUMMARY All the life history of the sea-slug Æolidiella alderi has been observed. The embryology, the metamorphosis and the post- larval development have been studied in detail from life and with the help of seriated and orientated histological sections of various stages from the gastrula to the adult. A. — This study shows especially : — Firstly, the structure of the nervous system and its evo­ lution. — Secondly, the reality of a 180° torsion. — Thirdly, in the inner part of the palliai fold a cellular proliferation arises, covering the visceral mass backward to give the definitive tegument and papillæ. — Fourthly, the realisation of a lengthened form, with a flat visceral mass closely linked with the cephalopodium shows the impossibility of an apparent complete detorsion as it is proved by normal and teratologic comparative morphogenesis. — Fifthly, the great similitarity of organisation in veligers with a protoconch of type I in Doridacea, Dendronotacea and Æolidiacea which contain a large amount of yolk in their eggs. MÉTAMORPHOSES DES NUDIBRANCHES B. — Comparison with the other kinds of metamorphosis in Nudibranches : I o Æolidiacea. I with planctotroph veligers. Some veligers crowling on their suitable prey have been obser­ ved just before they cast off their protoconch. These larval stages, which appeared to be those of Facelina coronata and of an undetermined species, show that the shell grows more than one turn of spire during their pelagic life. This last period is longer than that of veligers from Æolidiacea 2. 2° Æolidiacea 2 (with veligers of type 2 protoconch). In this case, however they are planctotrophic or lecithotrophic veligers, the part of the mantle fold which delimits the palliai cavity withdraws at hatching. So, the larval shell cannot enlarge during the pelagic phase. The notum and dorsal papil­ lae arise from the floor and the lateral parts of the mantle cavity. Æolidiacea 2 are more evolved and the pelagic stage is shorter than inÆolidiacea I. 3° Pleurobranchacea : Morphogenesis in Berthella plumula is more progressive than in unshelled Nudibranches. It arises according a very similar process to the one observed in Nudi­ branches I. C. — The discussion lies on : I o Torsion and its adaptative significance, 2° Chronology of metamorphosis process, 3° Teratology, natural and experimental morphogenesis in Nudibranches, 4° Nervous system (There is a true brain), 5° General phylogenetic conclusions; evolution within Nudi­ branches : a) From embryology : an early cleavage appears between Doridacea and other Nudibranches, but another one separates Nudibranches I and 2. These last ones would be more evolved. However, the Æolidiacea are probably not polyphyletic. b) From teratology and comparative anatomy : these facts show that branchiae is a true ctenidium inDoridacea, where all the steps from Pleurobranchacea to the most evolved Dori­ dacea can be found. We propose the sub-orderEuctenidiacea for the whole family Doridacea, which is opposed to the n. s/o Actenidiacea, covering all the other Nudibranches with secon­ dary or no branchial apparatus and a lateral anus. The Actenidiacea include the new superfamily Pseudoeucte­ nidiacea (for the genus Doridoxa which is not a Doridacea), and superfamilies Dendronotacea, Arminacea and Æolidiacea. The Nudibranches (includingPleurobranchacea) constitute a homogeneous whole from the evolutive point of view, and a clearly definite and natural order. 302 JEAN TARDY SOMMAIRE R é s u m é ........................................................................................................................................................................................................ 9 29 S u m m a r y ........................................................................................................................................................................................................................ 300 I ntroduction ............................................................................................................................................................................................................. 30 3 I. M o y e n s e t m é t h o d e s ................................................................................................................................................................................. 305 II. O n t o g e n è s e d ’Æ o l i d i e l l a a l d e r i (à 19-21° C)............................................................................... 30 7 1 . Clivage et gastrulation .............................................................................................................. 30 7 2 . Evolution de l’embryon jusqu’à la mise en place des ébauches oculaires.... 30 7 Stornodeum; vélum; pied; glande coquillière, manteau et bourrelet palléal; opercule; cellules anales; mésentéron; intestin; statocystes; « vésicules pulsátiles »; appareils excréteurs; muscles rétracteurs larvaires; tache violette; ganglions cérébro-pleuraux; ganglions buccaux; ganglions pédieux; commissures, connectifs et plaque sincipitale; ébauches oculaires. 3. Evolution de la véligère jusqu’à réversion complète du bourrelet palléal.... 316 Ganglions céphalo-pédieux; commissure viscérale; bulbe radulaire et arma­ ture buccale; mésentéron; intestin; cellules anales; protoconque; oper­ cule; « vésicules pulsátiles » ; canal réno-péricardique et cœur définitif; appareil excréteur; vélum; pied; gonade; manteau, formation de la cavité palléale; éversion; conséquences de Réversion : perte de la proto­ conque, position des orifices. 4 . Evolution jusqu’au stade pseudovermis ........................................................................... 3 2 3 Evolution morphologique : Achèvement du recouvrement de la masse viscérale par le tégument définitif et allongement. Conséquences de l’allongement : détorsion et réduction de la flexion ventrale; effacement de la masse viscérale; changements de position de certains organes. Evolution anatomique : Système nerveux : cérébralisation, télencéphalisa- tion; formation des ganglions rhinophoriens; innervation des palpes; ganglions pédieux; tube digestif : bulbe radulaire et glandes annexes; estomac et glandes digestives; vélum; rein; « vésicules pulsátiles »; cœur définitif; ébauche génitale; pied. 5 . Evolution
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